Effect of mutual shading on the emergence of nodal roots and the root/shoot ratio of maize

The effect of mutual shading on the root/shoot ratio and on the number of nodal roots of maize was studied. Plants of two varieties (Dea and LG2281) were grown in individual pots of 9 L, at three plant densities: 7.5, 11 and 15 plants m^–2. A control experiment was carried out in order to study if root growth was affected by the small size of the pots. Maize plants (cv Dea) were grown at a low plant density (7.5 plants m^–2) in pots of two different volumes (9 and 25 L respectively). In both experiments plants were watered every two hours with a nutrient solution. Some plants were sampled at five dates in the main experiment and the following data were recorded: foliar stage; root, stem and leaf dry weight; number of root primordia and number of emerged roots per phytomer. The final sampling date occurred at silking.Results of the control experiment showed that the root biomass was lower in small pots but the number of nodal roots per phytomer was not affected.Results of the main experiment showed that the total plant biomass and the root/shoot ratio were lower at high plant density. The number of emerged roots was strongly reduced on the upper phytomer (P₈). This reduction was mainly due to a lower percentage of root primordia which elongated. A proposed interpretation is that the number of roots which emerge on upper phytomers is controlled by carbohydrate availability.     

Plant Cd2+ and Zn2+ status effects on root and shoot heavy metal accumulation in Thlaspi caerulescens

* In this study we address the impact of changes in plant heavy metal, (i.e. zinc (Zn) and cadmium (Cd)) status on metal accumulation in the Zn/Cd hyperaccumulator, Thlaspi caerulescens. * Thlaspi caerulescens plants were grown hydroponically on both high and low Zn and Cd regimes and whole-shoot and -root metal accumulation, and root (109)Cd(2+) influx were determined. * High-Zn-grown (500 microm Zn) plants were found to be more Cd-tolerant than plants grown in standard Zn conditions (1 microm Zn). Furthermore, shoot Cd accumulation was significantly greater in the high-Zn-grown plants. A positive correlation was also found between shoot Zn accumulation and increased plant Cd status. Radiotracer (109)Cd root flux experiments demonstrated that high-Zn-grown plants maintained significantly higher root Cd(2+) influx than plants grown on 1 microm Zn. It was also found that both nickel (Ni) and copper (Cu) shoot accumulation were stimulated by high plant Zn status, while manganese (Mn) accumulation was not affected. * A speculative model is presented to explain these findings, suggesting that xylem loading may be one of the key sites responsible for the hyperaccumulation of Zn and Cd accumulation in Thlaspi caerulescens.     

Biomechanics of nodal, seminal and lateral roots of barley: effects of diameter, waterlogging and mechanical impedance

Background and aims

Biomechanical properties of cereal root systems largely control both resistance to root lodging and their ability to stabilise soil. Abiotic stresses can greatly modify root system growth and form. In this paper the effect of waterlogging and moderate mechanical impedance on root biomechanics is studied for both lateral roots and the main axes of barley.

Methods

Barley (Hordeum vulgare) plants were subjected to transient water-logging and moderate mechanical impedance in repacked soil columns. Roots were excavated, separated into types (nodal, seminal or lateral) and tested in tension to measure strength and elastic modulus.

Results

Water-logging and mechanical impedance substantially changed root system growth whilst root biomechanical properties were affected by waterlogging. Root strength was generally greater in thin roots and depended on root type. For example, seminal roots 0.4–0.6 mm in diameter were approximately seven times stronger and five times stiffer than lateral roots of the same diameter when mechanically impeded. Root sample populations typically exhibited negative power-law relationships between root strength and diameter for all root types. Mechanical impedance slowed seminal root elongation by approximately 50 % and resulted in a 15 % and 11 % increase in the diameter of in nodal and seminal roots respectively. Power-law relationships between root diameter and root biomechanical properties corresponded to the different root types. Coefficients for between root diameter, strength and elastic modulus improved when separated by root type, with R² values increasing in some roots from 0.05 to 0.71 for root strength and 0.08 to 0.74 for elastic modulus.

Conclusions

Moderate mechanical impedance did not influence the tensile strength of roots, but, waterlogging diminished the relationship between root strength and diameter. Separation of root type improved predictions of root strength and elastic modulus using power-law regressions.     

Species-specific effects of live roots and shoot litter on soil decomposer abundances do not forecast plant litter-nitrogen uptake

Plant species produce litter of varying quality and differ in the quality and quantity of compounds they release from live roots, which both can induce different decomposer growth in the soil. To test whether differences in decomposer growth can forecast the amount of N species acquire from plant litter, as suggested by theory, we grew individuals of three grassland plants—Holcus lanatus, Plantago lanceolata and Lotus corniculatus—in soils into which ¹⁵N-labelled litter of either Holcus, Plantago or Lotus was added. We measured the effects of live roots and litter of each species on soil microbes and their protozoan and nematode feeders, and to link decomposer growth and plant nutrient uptake, we measured the amount of N taken up by plants from the added litter. We hypothesised that those species that induce the highest growth of microbes, and especially that of microbial feeders, will also take up the highest amount of N from the litter. We found, however, that although numbers of bacterial-feeding Protozoa and nematodes were on average lower after addition of Holcus than Plantago or Lotus litter, N uptake was higher from Holcus litter. Further, although the effects on Protozoa and bacterial- and fungal-feeding nematodes did not differ between the live plants, litter-N uptake differed, with Holcus being the most efficient compared to Plantago and Lotus. Hence, although microbes and their feeders unquestionably control N mineralization in the soil, and their growth differs among plant species, these differences cannot predict differences in litter-N uptake among plant species. A likely reason is that for nutrient uptake, other species-specific plant traits, such as litter chemistry, root proliferation ability and competitiveness for soil N, override in significance the species-specific ability of plants to induce decomposer growth.     

The effects of stretch activation on ionic selectivity of the TREK-2 K2P K+ channel

The TREK-2 (KCNK10) K2P potassium channel can be regulated by variety of polymodal stimuli including pressure. In a recent study, we demonstrated that this mechanosensitive K⁺ channel responds to changes in membrane tension by undergoing a major structural change from its ‘down’ state to the more expanded ‘up’ state conformation. These changes are mostly restricted to the lower part of the protein within the bilayer, but are allosterically coupled to the primary gating mechanism located within the selectivity filter. However, any such structural changes within the filter also have the potential to alter ionic selectivity and there are reports that some K2Ps, including TREK channels, exhibit a dynamic ionic selectivity. In this addendum to our previous study we have therefore examined whether the selectivity of TREK-2 is altered by stretch activation. Our results reveal that the filter remains stable and highly selective for K⁺ over Na⁺ during stretch activation, and that permeability to a range of other cations (Rb⁺, Cs⁺ and NH₄⁺) also does not change. The asymmetric structural changes that occur during stretch activation therefore allow the channel to respond to changes in membrane tension without a loss of K⁺ selectivity.     

Causes and location of non-specific effects of SHAM on O2 uptake by wheat roots

Uptake of O₂ by whole, detached, root systems of wheat ( Triticum aestivum L. cv. Alexandria) was titrated with salicylhydroxamic acid (SHAM) in the presence and absence of cyanide. The resulting Q_all plot was non-linear indicating that SHAM was acting non-specifically. The nature of the non-specific effects was investigated in reverse titration experiments. Uptake of O₂ was titrated with KCN in the presence and absence of SHAM at 1 m M and 25 m M , which yielded Q_cy1 values of < 1 and > 1, respectively. The results suggest that at 25 m M , SHAM inhibits the cytochrome pathway, but at 1 m M it stimulates an O₂-consuming process which is likely to be a peroxidase. A SHAM-stimulated peroxidase could easily be washed from these roots. In vitro, the peroxidase was stimulated to a similar extent by low (1 m M ) and high (25 m M ) concentrations of SHAM. Failure to inhibit with high concentrations of SHAM distinguishes this peroxidase from those bitherto eluted from root tissue. Reverse titration experiments in the presence and absence of 1 m M SHAM indicated that there were no significant side effects of SHAM in root tips. These data are supported by the negligible peroxidase activity that was washed from this root fraction. In contrast, significant side effects occurred in vivo, and substantial peroxidase activity was measured in vitro, from sections 4–6 cm and 18–20 cm behind the seminal root apex. The greatest activity was found with the 4–6 cm section which may be associated with high rates of cell wall lignification. The implications of these results for measurements of root respiration are discussed.     

Nitrite in the root zone and its effects on ion uptake and growth of wheat seedlings

The immediate and posteffects of various concentrations of NaNO₂ on ion uptake of wheat ( Triticum aestivum L. cv. GK Öthalom) seedlings were studied at different pH values. Without pretreatment, the higher the concentration of NaNO₂ the greater was the decrease in uptake of K⁺ into the roots, both at pH 4 and pH 6. At pH 6 but not at pH 4 the reverse was true when the seedlings were pretreated with NaNO₂. Due to the high Na⁺ content of the roots, an effect of Na⁺ in this process cannot be excluded. Nitrite was taken up by the roots more rapidly than nitrate. Nitrite at 0.1 m M in the medium induced the development of an uptake system for both NO⁻₂ and NO⁻₃ in wheat roots. At higher concentrations pretreatment with NO⁻₂ decreased NO⁻₃ uptake by the roots, but NO₃ did not inhibit the uptake of NO₂. The toxic effect of NO⁻₂ was strongly pH dependent. Lower pH of the external solution led to an increased inhibition by NO⁻₂ of both ion uptake and growth of seedlings. The inhibitory effect of NO⁻₂ differed considerably for roots and shoots. The roots and especially the root hairs were particularly sensitive to NO⁻₂ treatment.     

Soil O2 and CO2 effects on root respiration of cacti

Through use of a recently developed technique that can measure CO₂ exchange by individual attached roots, the influences of soil O₂ and CO₂ concentrations on root respiration were determined for two species of shallow-rooted cacti that typically occur in porous, well-drained soils. Although soil O₂ concentrations in the rooting zone in the field were indistinguishable from that in the ambient air (21% by volume), the CO₂ concentrations 10 cm below the soil surface averaged 540 μLL⁻¹ for the barrel cactusFerocactus acanthodes under dry conditions and 2400 μLL⁻¹ under wet conditions in a loamy sand. For the widely cultivated platyopuntiaOpuntia ficus-indica in a sandy clay loam, the CO₂ concentration at 10 cm averaged 1080 μLL⁻¹ under dry conditions and 4170 μLL⁻¹ under wet conditions. For both species, the respiration rate in the laboratory was zero at 0% O₂ and increased to its maximum value at 5% O₂ for rain roots (roots induced by watering) and 16% O₂ for established roots. Established roots ofO. ficus-indica were slightly more tolerant of elevated CO₂ than were those ofF. acanthodes, 5000 μLL⁻¹ inhibiting respiration by 35% and 46%, respectively. For both species, root respiration was reduced to zero at 20,000 μLL⁻¹ (2%) CO₂. In contrast to the reversible effects of 0% O₂, inhibition by 2% CO₂ was irreversible and led to the death of cortical cells in established roots in 6 h. Although the restriction of various cacti and other CAM plants to porous soils has generally been attributed to their requirement for high O₂ concentrations, the present results indicate that susceptibility of root respiration to elevated soil CO₂ concentrations may be more important.     

Water uptake by roots of Hordeum marinum: formation of a barrier to radial O2 loss does not affect root hydraulic conductivity

The adventitious roots of Hordeum marinum grown in stagnant deoxygenated solution contain a barrier to radial O2 loss (ROL) in basal zones, whereas roots of plants grown in aerated solution do not. The present experiments assessed whether induction of the barrier to ROL influences root hydraulic conductivity (Lpr). Wheat (Triticum aestivum) was also studied since, like H. marinum, this species forms aerenchyma in stagnant conditions, but does not form a barrier to ROL. Plants were grown in either aerated or stagnant, deoxygenated nutrient solution for 21-28 d. Root-sleeving O2 electrodes were used to assess patterns of ROL along adventitious roots, and a root-pressure probe and a pressure chamber to measure Lpr for individual adventitious roots and whole root systems, respectively. Lpr, measured under a hydrostatic pressure gradient, was 1.8-fold higher for individual roots, and 5.6-fold higher for whole roots systems, in T. aestivum than H. marinum. However, there was no difference in Lpr between the two species when measured under an osmotic driving force, when water moved from cell to cell rather than apoplastically. Root-zone O2 treatments during growth had no effect on Lpr for either species (measured in aerobic solution). It is concluded that induction of the barrier to ROL in H. marinum did not significantly affect the hydraulic conductivity of either individual adventitious roots or of the whole root system.     

水磷耦合对小麦次生根特殊根毛形态与结构的影响

通过水、磷复因子大田试验,以强筋小麦品种郑麦9023为材料,研究了水磷耦合对小麦生育中、后期次生根特殊根毛形态与结构的影响。结果指出,不同水分处理显著影响特殊根毛形态。随着土壤水分含量提高,次生根特殊根毛长度缩短。与土壤湿润处理相比,仅依靠自然降水处理的特殊根毛长度和直径增加(P<0.01),拔节至子粒形成期间完全灌溉处理的根毛长度增加(P<0.01)。随着供磷水平提高,特殊根毛长度和直径增加(P<0.05),其中高磷处理和对照（不施磷）的差异达极显著水平(P<0.01)。同一供水条件下随供磷水平提高,或同一供磷水平上随土壤含水量降低,特殊根毛长度和直径均增加(P<0.05)。拔节以后,仅依靠自然降水—高磷处理组合的特殊根毛细胞饱满,结构完整,细胞壁加厚明显,细胞核、液泡及线粒体清晰可见;而土壤湿润—低磷处理组合的特殊根毛扭曲、变形现象严重,细胞壁变薄,细胞核解体,质膜、微体等细胞器消失。研究表明,适当降低土壤含水量并提高供磷水平,小麦次生根特殊根毛的长度和直径增加,并维持良好的细胞形态和结构。     

Effects of salinity on uptake and distribution of Na+, Cl- and K+ in two wheat cultivars

Plants of two wheat (Triticum aestivum L.) cultivars differing in salt tolerance were grown in sand with nutrient solutions. 35-d-old plants were subjected to 5 levels of salinity created by adding NaCl, CaCl₂ and Na₂SO₄. Growth reduction caused by salinity was accompanied by increased Na⁺ and Cl^- concentrations, Na⁺/K⁺ ratio, and decreased concentration of K⁺. The salt tolerant cv. Kharchia 65 showed better ionic regulation. Salinity up to 15.7 dS m^-1 induced increased uptake of Na⁺ and Cl^- but higher levels of salinity were not accompanied by further increase in uptake of these ions. Observed increases in Na⁺ and Cl^- concentrations at higher salinities seemed to be the consequence of reduction in growth. Uptake of K⁺ was decreased; more in salt sensitive cultivar. This was also accompanied by differences in its distribution.     

O3 impacts on plant development: a meta-analysis of root/shoot allocation and growth

The mechanism of O3 action on plants remains poorly characterized. Symptoms include visible lesions on the leaf surface, reduced growth and a hypothesized reduction in allocation of carbohydrate to roots. The generality of this latter phenomenon has not been demonstrated. Here, a meta-analysis is performed of all available experimental data, to test the hypotheses that O3 exposure of the shoot inhibits biomass allocation below ground (the root/shoot allometric coefficient, k) and inhibits whole-plant growth rate [relative growth rate (RGR)]. Both k and RGR were significantly reduced by O3 (5.6 and 8.2%, respectively). Variability in k was greater than in RGR, and both exhibited some positive as well as mostly negative responses. The effects on k were distinct from the effects on RGR. In some cases, k was reduced while RGR was unaffected. Slow-growing plants (small RGR) exhibited the largest declines in k. These observations may have mechanistic implications regarding O3 phytotoxicity. There were no effects of type of exposure chamber on sensitivity to O3. The analyses indicate that the O3 inhibition of allocation to roots is real and general, but variable. Further experiments are needed for under-represented plant groups, to characterize exceptions to this generalization and to evaluate O3--environment interactions.     

Environmental effects on dry matter partitioning between shoot and root of crop plants: relations with growth and shoot protein concentration

The literature on environmental effects on dry matter partitioning in higher plants, in particular crop plants, is reviewed focussing on changes in shoot to root dry weight ratio (S:R). Of particular consistency is the finding that S:R increases with increased nitrogen (N) supply. Relations between nitrogen (N) supply, growth, S:R and tissue N and protein concentration are examined. In some cases, the increase in S:R with increased N supply is likely to have been at least in part an effect on growth and development, but there is unequivocal evidence that N affects S:R independently of growth and development. A positive correlation between S:R and leaf protein concentration is highlighted. It is argued that the N effect on S:R outside the effect on growth and development is related to increased shoot protein concentration. Specifically, shoot and root growth are colimited by local carbon (C) and N (primarily protein) substrate concentrations and shoot growth will increase relative to root growth with increased N substrate availability due to the proximity of the shoot to the C source. It is further argued that results in the literature are consistent with the proposal that macronutrient, water, irradiance, CO₂ and temperature effects on S:R are often primarily mediated through their effects on growth and development, and shoot protein concentration and hence shoot growth.     

Effect of Fe2+, Mn2+, Zn2+ and Pb2+ on H+/K+ fluxes in excisedPistia stratiotes roots

Pistia stratiotes is used for the epuration of domestic sewage in the Biyem Assi phytopurification station. During the process, Fe²⁺, Mn²⁺, Zn²⁺ and Pb²⁺ are absorbed in substantial amounts by the plant. These metals modify the H⁺/K⁺ exchange system at the root level. H⁺ efflux is inhibited by Fe²⁺ and by Zn²⁺ and enhanced by Mn²⁺ and Pb²⁺. K⁺ influx is inhibited by Fe²⁺, by Zn²⁺ and by Pb²⁺ and enhanced by Mn²⁺. It is shown that the purification capacity ofPistia stratiotes can vary with the composition of the heavy metals in the surrounding medium.     

Control of root growth: effects of carbohydrates on the extension, branching and rate of respiration of different fractions of wheat roots

Rates of extension, numbers of laterals and rates of respiration were measured in different fractions of wheat ( Triticum aestivum L. cv. Alexandria) roots following changes in carbohydrate supply. The supply of carbohydrate was varied by selective pruning and exogenously fed sugars. Pruning shoots to a single leaf (leaf-pruning) reduced the rate of O₂ uptake by intact roots. Rates were not stimulated by shortterm feeding of sucrose (25 m M ), but were stimulated by the uncoupler p -trifluoro-methoxy(carbonylcyanide)phenylhydrazone (FCCP). Feeding glucose to roots of leaf-pruned and non-pruned plants for 16–24 h increased the rate of O₂ uptake. It is concluded that respiration is under fine control by adenylates and coarse control by carbohydrate supply, with carbohydrates regulating directly the rate of some energy consuming process(es). These energy consuming processes are located in growing tissue fractions. Feeding glucose to leaf-pruned and non-pruned plants increased rates of O₂ uptake in seminal root tips, the zone of developing lateral primordia and mature root sections with elongating laterals, but had no effect on mature sections from which the laterals had been excised. Leaf-pruning reduced the extension rate of seminal axes and first-order laterals when measured over 24 h. Feeding glucose to roots from the time of pruning increased the rate, but did not fully restore it to control values. Pruning roots to a single seminal axis (root-pruning) and feeding glucose to non-pruned plants had no effect on the extension rate of the seminal axis or its laterals over this time period, although rates were increased by root-pruning when measured over 3 days. The number of lateral root primordia was reduced by leaf-pruning and increased by root-pruning and feeding glucose. The results are discussed in terms of the role of carbohydrates in the control of root growth and branching.     

冬小麦生育中后期次生根特殊根毛的发生与形态结构

大田试验条件下,研究了冬小麦次生根特殊根毛的发生规律和形态结构特征.结果表明,拔节后次生根近植株基部根体上特殊根毛普遍发生,其在次生根根体上的分布可区分为集中区、适中区和稀少区.特殊根毛集中区根毛的长度、直径、密度分别均大于适中区和稀少区,其中,长度和密度的差异达极显著水平(P<0.01).随着生育期推进, 特殊根毛长度、直径和密度均呈下降趋势.大多数特殊根毛呈现出不同程度的扭曲、变形, 根毛细胞突起和分枝现象较为普遍,特殊根毛细胞次生壁出现加厚现象.     

Ionic mechanisms of action of GABA on dorsal and ventral root myelinated fibers: effects of K+ channel blockers

Excitability changes evoked by the inhibitory neurotransmitter, GABA (gamma-aminobutyric acid) in myelinated axons of dorsal and ventral roots of the isolated bullfrog sciatic nerve were compared in the absence and presence of K+ channel blockers. Half-maximal A-fiber responses to a 0.5-Hz stimulation of the whole nerve were recorded from individual roots. Direct applications of Ringer with raised K+ levels to the site of stimulation caused increases in excitability of both dorsal and ventral root fibers, which resembled those evoked in the ventral root by the GABA agonist THIP (4,5,6,7-tetrahydroisoxazolo[5,4-c]ol). The increases in dorsal root fiber responses produced by GABA were depressed by tetraethylammonium (TEA) (3 mM), 4-aminopyridine (4-AP) (50 microM), Cs (2 mM), and Ba (1 mM). Ventral root fibers were less consistently affected. The early component of GABA-evoked excitability increases was depressed by 4-AP, Cs, and Ba, but greatly augmented by TEA. THIP-evoked changes in the excitability of the dorsal and ventral root fibers were, respectively, depressed and enhanced by TEA. The augmenting effect of TEA on the early component of GABA agonist effects on the ventral root fibers is attributed to their high resting K+ conductance and the presence of a slowly inactivating, fast K+ current (If1). The depressant effects of K+ channel blockade on depolarizing components of agonist-evoked changes in dorsal and ventral root responses indicate interference with release and (or) sensitivity to K+ and a possible contribution from a mechanism involving voltage-dependent delayed rectifier K+ currents.